Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene

Quaternary International xxx (2015) 1e11

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Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene P.A. Kosintsev*, V.V. Gasilin, D.O. Gimranov, O.P. Bachura Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences, Russian Federation

a r t i c l e i n f o

a b s t r a c t

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Carnivore assemblages from localities of the Late Pleistocene and Holocene of Northern, Middle and Southern Urals were studied. Analysis of species composition of the carnivore fauna was performed for seven chronological periods: MIS 5e, MIS 3, mid-MIS2 (LGT), late MIS 2 (BAIC), early MIS 1 (PreborealBoreal), mid-MIS 1 (Atlantic-Subboreal), late MIS 1 (Subatlantic). In the Urals, changes in carnivore fauna were occurring during the Late Pleistocene and Holocene. Faunal composition included the species from European, Siberian and EuropeaneSiberian faunal complexes throughout the Late Pleistocene and Holocene, dominated by the species from EuropeaneSiberian faunal complexes. The general trend of fauna changes is represented by a decreasing percentage of species from the European faunal complex and an increasing percentage of those from the Siberian faunal complex. Faunal composition has changed due to two main reasons, range shifts and extinction. Sixteen out of 24 species that inhabited the Urals during the Late Pleistocene and Holocene underwent range shifts. The ranges shifted in different directions: reduction, increased and fluctuations. The ranges of several species have not changed. Three species became extinct. © 2015 Elsevier Ltd and INQUA.

Keywords: Carnivora Ural Late Pleistocene Holocene Fauna Range

1. Introduction The Ural region has overlapping ranges of species from the different geographical fauna complexes: European, Siberian, and Euro-Siberian (Kuznetsov, 1950; Kulik, 1980). It is located on the border between Europe and Asia, and crosses tundra, taiga, and steppe natural zones. At present, the Urals is inhabited by 16 species of carnivores belonging to four families (Canidae, Ursidae, Mustelidae, and Felidae) (Marvin, 1969). The Urals is known for its well-studied Late Pleistocene and Holocene localities, yielding numerous fossil mammal assemblages, including fossil carnivores. The results of these studies have been presented in a considerable number of papers dedicated either to the description of the history of large mammals of selected areas of the Urals (Kuzmina, 1971, 1975; Smirnov et al., 1990; Kosintsev and Bachura, 2005, 2013, 2014; Ulitko, 2006; Bachura and Kosintsev, 2007; Kosintsev, 2007; Kosintsev and Plasteeva, 2009; Danukalova et al., 2011; Ponomarev et al., 2013; Smirnov et al., 2014) or to the description of the fauna of certain localities (Bachura and Strukova, 2002; Bachura and Plasteeva, 2005; Gimranov, 2009; Kosintsev, 1996; Kosintsev and Orlova, 2002; Petrov, Kosintsev, 2005; Kuzmina et al., 1999; Kuz'mina, 2000;

Razhev et al., 2005; Strukova et al., 2006; Svendsen et al., 2010; Ulitko et al., 2011; Yakovlev et al., 2006). In these works, carnivores are studied together with other large mammals. Some articles devoted to the history of certain carnivore species of the Urals have been published (Baryshnikov, 2001, 2007; Gasilin and Kosintsev, 2010; Gasilin et al., 2014; Gimranov and Kosintsev, 2015; Kosintsev and Bachura, 2015; Kosintsev and Gasilin, 2011; Kosintsev, Vorob'ev, 2001; Kuzmina, 2002). The purpose of this paper is to reconstruct the history of carnivore fauna of the Urals in the Late Pleistocene and Holocene. The most ancient carnivore fossils have been found in the Southern Urals. At the site of Baturino, remains of Ursus sp. and Mustela (small forma) whose size is close to that of M. erminea and M. nivalis, have been found in the deposits dating to the middle stages of the Early Pleistocene, while the Mustela remains (small form) date to the end of the Early Pleistocene (Stefanosvkiy and Borodin, 2002). The Middle Pleistocene deposits of the Ignatievskay cave contain remains of Canis lupus and Canis sp. (small form), Vulpes vulpes, V. lagopus, Ursus savini, Martes sp., Gulo gulo, Mustela erminea, and M. nivalis (Smirnov et al., 1990). The Late Pleistocene deposits yield other Uralian carnivore remains. 2. The modern situation: some components of the Urals biota

* Corresponding author. E-mail address: [email protected] (P.A. Kosintsev).

Four regions are traditionally distinguished within the Urals: the Polar (from 67
300 N to 64
000 N), Northern (from 64
000 N to

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Please cite this article in press as: Kosintsev, P.A., et al., Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.10.089

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P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

59
150 N), Middle (from 59
150 N to 55
540 N), and Southern Urals (from 55
540 N to 51
000 N) (Urals and Pre-Urals, 1968). Although this division is based on the orography of the Ural Range, each of these regions has its specific natural and climatic features. Here, we consider the Middle and Southern Urals in a broad sense, including the mountains and parts of the Trans-Urals peneplain. We do not have data on the carnivore fauna history in the Polar Urals, so we do not consider this region in this article. The climate of the Northern, Middle, and Southern Urals is temperate continental. The Urals is disposed across the direction of the prevailing westerly winds, so this western slope is moister than the east slope. The territories of Northern and Middle Urals are located in the taiga where pine and spruce dominate. In the south part of the Middle Urals, there is a large proportion of birch in the composition of vegetation. Almost all the mountainous part of the Southern Ural territory lies in the zone of coniferous small-leaf and coniferous broadleaf forests. Pine, birch, linden, elm and oak (in the south) dominate among vegetation. The southern part of the Southern Urals is located in the steppe zone. In the peneplain of the Middle and Southern Urals, steppe vegetation is present. The mountains of the Northern, Middle and Southern Urals are not high so the altitudinal belt in vegetation is not expressed (Gorchakovskiy, 1975). Fauna of carnivore of the Urals consist of three main complexes: tundra, forest, and steppe. The carnivores which inhabit the entire territory of Urals are Canis lupus, Vulpes vulpes, M. nivalis, M. erminea. A typical tundra species is Vulpes lagopus which inhabits the Polar Urals and the extremely northern part of the Northern Urals. The forest complex of carnivores inhabits territory of Northern and Middle Urals as well as mountain-forest zone of Southern Urals. This complex is represented by the taiga (Ursus arctos, Gulo gulo, Martes zibellina, Martes martes, M. putorius, M. sibirica, Lynx lynx) and taiga-wetland species (M. lutreola, M. vison, Lutra lutra). The species of forest-steppe species (Meles leucurus) inhabits the southern Northern Urals, the Middle, and Southern Urals. Distribution of carnivores of the steppe mammal complex is limited by the southern part of the Middle Urals and Southern Urals (M. eversmanni) and the Southern Trans-Urals peneplain area (V. corsac) (Urals and Pre-Urals, 1968). 3. Materials and methods We studied data on carnivore species composition from 287 sites located (Table A.1.) in the Urals between 64
and 51
N (Fig. 1). The carnivore remains have been found in localities of several types, i.e. alluvial, swamp, cave, and archaeological localities. All faunas have been dated. The MIS 5 faunas have been dated on the basis of the species composition of the associated mammal fauna. All the MIS 3 and MIS 2 faunas have been dated using radiocarbon. The MIS 1 fauna has been dated using radiocarbon and archaeological methods. Radiocarbon dates from the Uralian localities were obtained from the bones of several carnivore species (Table A.2.). We use uncalibrated radiocarbon dates. All radiocarbon dates with indices GIN, LE, LU, SOAN were obtained by conventional radiocarbon dating, while the dates with other indices by accelerator mass spectrometer (AMS) method. Some localities were radiocarbon-dated by both methods. The comparison of radiocarbon dates obtained by different methods for the cave bear bones indicates that sometimes these methods do not give consistent results (Table A.2.). For example, a conventional radiocarbon date of 26980 BP and some AMS dates, ranging between 37190 BP and 47600 BP, were obtained for the Tayn cave. The bones from the Geologov 3 cave were a source for two conventional radiocarbon dates of 27070 BP and 27580 BP, as well as a single AMS date of 38480 BP. The bones from the Zapovednaya cave gave

two conventional dates of 28700 BP and 37250 BP and three infinite AMS dates. The general trend is that the conventional 14C dates (i.e., produced by liquid scintillation counting) yield older ages compared to the AMS 14C values from the same strata. Therefore, we will use AMS dates to determine the dates of species extinction. The available data allowed us to perform the analysis of the species composition of the carnivore fauna for seven chronological periods: MIS 5e, MIS 3, mid-MIS 2 (LGT), late MIS2 (BAIC), early MIS 1 (Preboreal-Boreal, Early Holocene), mid-MIS 1 (Atlantic-Subboreal, Middle Holocene), late MIS 1 (Subatlantic, Late Holocene). Data for the other periods are not sufficient. In the Late Interglacial (MIS 5e, Eemian, Mikulino, Streletsk, Kazantsevo), the climate was warmer than now, and forest vegetation was more widespread (Mangerud, 1989; Grichuk, 2002; Kühl, Litt, 2003). The mega-interstadial of the Last Glaciation (MIS 3, Middle Weichselian, Bryansk Interstadial, Karginsk Interstadial) with an age of 60e24 ka BP is characterized by relatively warm climate but also by the occurrence of a large number of climatic fluctuations which differ in intensity (Zagwijn, 1974; Vanderberge, 2002; Rasmussen et al., 2006; Svensson et al., 2008; Velichko and Faustova, 2009). As for the end of the Late Pleistocene (MIS 2, Late Weichselian, Late Valdai, Sartan), we have studied two periods that occurred after the Last Glacial Maximum (LGM), i.e. Deglaciation or Late Glacial Transition (LGT; 17e12.4 ka BP) and the Bølling e Allerød interstadial warming divided by short and weak pronounced Older Dryas Stadial (BAIC; Late Weichselian, Late Valdai, Late Sartan; 12.4e10.3 ka BP) (Markova et al., 2008; Faustova, 1994). At the onset of the Holocene, the climate became warmer (Svensson et al., 2008) and the forests spread rapidly (Chotinsky and Klimanov, 2002). The chronology of the localities examined in the paper covers the entire the Late Pleistocene and Holocene, from MIS 5e to late MIS 1. The boundaries of the chronological periods are given as radiocarbon years. The faunas of separate localities within one area (Northern, Middle and Southern Urals) have been chronologically classified into several groups (Table A.1.). The Southern Urals includes two large geomorphologic units: the mountain region and the Transuralian peneplain. Within the territory of the Southern Urals, all the Late Pleistocene localities are situated in mountains region. Therefore, we consider them as one geographical group. The Holocene localities are situated in both regions. Thus, the localities of each Holocene period form two geographical groups, namely the “mountain” and the “peneplain” groups. The analysis of the Late Pleistocene and Holocene and modern ranges of each species allowed us to assign all the carnivore species to one of three faunal complexes, European, Siberian, or EuropeaneSiberian complex. The ranges of many species cover the territory of both Europe and Asia. The European faunal complex is represented by the species whose range lies mostly in Europe, and its eastern margin in Asia is restricted to the Yenisei River. The Siberian faunal complex contains the species whose range lies mostly in Asia and its western margin in Europe is restricted to the Volga basin. The EuropeaneSiberian faunal complex includes the species with their ranges represented mostly in both Europe and Asia. It is difficult to assign the Asiatic black bear (Ursus tibethanus) and dhole (Cuon alpinus) to a particular faunal complex. At present, their ranges lie entirely in Asia, though their Pleistocene ranges were different. During the Saalian glaciation the Asiatic black bear's range covered central (Musil, 2005e2006; Nagel, Rabeder, 2000) and southern Europe and extended into the Iberian Peninsula
gut-Bonnoure, 1997). This range apparently (Torres, 1988; Cre remained unchanged until the end of the Eem interglacial and was drastically reduced at the onset of the Weichselian Stage. The dhole's range covered southern and central Europe in the Late Pleistocene (Sommer and Benecke, 2005). The dhole persisted in

Please cite this article in press as: Kosintsev, P.A., et al., Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.10.089

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Fig. 1. The main localities of carnivore remains from the Urals. 1 e Medvezhaya cave; 2 e Kaninskaya cave; 3 e Uninskaya cave; 4 e Burmantovo; 5 e Usoltsevskaya cave; 6 e Kamen’ Pysannyj; 7 e Cheremukhovo-1; 8 e Zhilische Sokola; 9 e Makhnevskaya-Ledyanaya cave; 10 e Tayn cave; 11 e Kizelovskaya cave; 12 e Rasik; 13 e Verkhnegubakhinskaya cave; 14 e Lun'evka III cave; 15 e Viasher cave; 16 e Lobvinskaya cave; 17 e Grotto Bolshoj Glukhoj; 18 e Dyrovatyj-na-Chusovoj; 19 e Ust’-Katavskaya cave; 20 e Turistov cave; 21 e Koksharovsko-Ur'inskaya; 22 e Shigirskoe gorodische; 23 e Georgievskaya cave; 24 e Pershinskaya cave; 25 e Bezymaynnyj; 26 e Zotinskij; 27 e Grotto Bobylek; 28 e Barsuchij Dol; 29 e Ust’-Katavskaya cave; 30 e Ignat'evskaya cave; 31 e Pymva-Shor; 32 e Mamontova Kyr'ya; 33 e Pizhma-1; 34 e Pobeda cave; 35 e Nikol'skaya cave; 36 e Sikiyaz-Tamak; 37 e Bayslan-Tash; 38 e Balatukaj; 39 e Ustinovo; 40 e Zapovednaya cave; 41 e Imanaj cave; 42 e Verkhnyaya cave; 43 e Asha-1; 44 e Smelovskaya cave; 45 e Syrtinskaya cave; 46 e Severnaya cave; 47 e Laksejskaya cave; 48 e Shajtanskaya cave.

the Iberian Peninsula until the Early Holocene (Ripol et al., 2010). Thus, EuropeaneSiberian geographical distribution was characteristic of both Asiatic black bear and dhole in the Pleistocene. That is why we consider them to be part of the EuropeaneSiberian faunal complex. The ranges of several species with quite similar skeleton structure overlap or used to overlap each other in the Urals. We applied special methods for species identification of fossil remains of the following species: Vulpes vulpes, V. lagopus and V. corsac (Monchot and Gendron, 2010), Canis lupus and Cuon alpinus (Ripol et al., 2010; Pionnier-Capitan et al., 2011), Meles meles and M. leucurus (Aristov and Baryshnikov, 2001; Gasilin and Kosintsev, 2012), Martes martes, M. zibellina and M. foina (Gasilin and Kosintsev, 2013; Gimranov, 2015; Gimranov and Kosintsev, 2015). The analysis of ancient DNA from the cave bear bones from the Northern Urals showed that the Ursus (Spelaearctos) ingressus form

inhabited this area (Knapp et al., 2009). No DNA analysis has been performed for the bears from other Uralian areas. Pending data for other areas, we assign the cave bear of the Urals to Ursus (Spelaearctos) spelaeus. The approach representing order Carnivora follows Wilson and Reeder (2005). 4. Discussion The most ancient Late Pleistocene carnivore remains have been found in the faunas of the Bobylek caves (Smirnov, 1993; Razhev et al., 2005) and the Makhnevskaya Ledyanaya in the Middle Urals (Barysnikov, 2001; Fadeeva et al., 2011), as well as in the Barsuchii Dol in the Southern Urals (Fig. 1). As these faunas comprise forest species (Lynx lynx, Ursus tibethanus, Sciurus vulgaris, Apodemus flavicollis, Dryomys nitedula) and those that inhabit the areas with relatively warm climate (Desmana moshata,

Please cite this article in press as: Kosintsev, P.A., et al., Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.10.089

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Talpa europaea, Lutra lutra, Meles meles, Hystrix brachyura), they have been assigned to an interglacial time. Simultaneously the typical Late Pleistocene species were found with these species as part of the fauna e Ursus spelaeus, U. savini, Panthera spelaea, Mammuthus primigenius, Coelodonta antiquitatis, Bison priscus, Microtus gregalis and Lemmus sibiricus (Baryshnikov, 2001; Razhev et al., 2005; Fadeeva et al., 2011). This evidence allows us to allocate this interglacial to the Late Pleistocene and to date the faunas to Streletsk interglacial (MIS 5e, Eemian, Mikulino, Kazantsevo). Species composition of the Middle and Southern Uralian faunas was nearly the same at that time (Table 1). Asiatic black bear (Ursus tibethanus) (Barysnikov, 2001) and wolverine (Gulo gulo) (Razev et al., 2005) were present in the interglacial time fauna of the Middle Urals. The absence of these species in fauna of the Southern Urals can be explained by insufficient numbers of well-studied localities of this period. This might also explain the absence of some species of the genera Martes and Mustela in the composition of these faunas. The faunal composition is dominated by the species of the EuropeaneSiberian faunal complex. The species of the European faunal complex are not abundant, and the species of the Siberian faunal complex are absent (Table 2). The presence of the Asian black bear in the fauna of this period is regarded as its peculiarity (Table 1), as this species is absent from later faunas. Meles meles was part of the faunal composition of this period, though it is absent from the fauna of the subsequent periods.

Faunas from some dozens of the localities in the Northern, Middle and Southern Urals are known from the mega-interstadial of the Last Glaciation (MIS 3). These faunas comprised the species of the EuropeaneSiberian, European and Siberian faunal complexes (Table 1). All these faunas were dominated by the species of the EuropeaneSiberian faunal complex, while European and Siberian complexes were represented by rare species (Table 2). This period is characterized by differences in the species composition of the fauna typical of different areas. The number of species decreased from south to north. Eighteen species inhabited the Southern Urals, 14 species were common for the Middle Urals and 12 species occurred in the Northern Urals. The fauna of the Southern Urals was distinguished by the dhole (Cuon alpinus), lynx (Lynx lynx), otter (Lutra lutra), pine marten (Martes martes) and European mink (Mustela lutreola). These species were absent from the Middle Uralian fauna (Table 1). As for the Northern Urals, besides the above listed species, сave hyena (Crocuta crocuta spelaea) and corsac (Vulpes corsac) were also absent from its fauna (Table 1). In contrast to the fauna of the Southern Urals, faunal composition of the Middle and Northern Urals was distinguished by the sable (Martes zibellina). Forest (lynx, and semi-aquatic (otter, European mink) species were the first to disappear from the fauna from south to north, followed by relatively “thermophilic” species (corsaс, cave hyena) in passing northwards. The faunal composition included several species (Cuon alpinus, Crocuta crocuta spelaea, Ursus spelaeus, Ursus savini) which were absent from later periods. The ranges of the first two species

Table 1 Species composition of carnivore fauna from the Ural during the Late Pleistocene Species

Faunal complex

Lynx lynx Panthera spelaea Crocuta crocuta spelaea Canis lupus Cuon alpinus Vulpes corsac Vulpes lagopus Vulpes vulpes Ursus arctos Ursus tibethanus Ursus spelaeus Ursus savini Lutra lutra Gulo gulo Martes martes Martes zibellina Meles leucurus Meles meles Mustela erminea Mustela eversmanii Mustela lutreola Mustela nivalis a

MIS 5е

EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian Siberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian European EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian European Siberian Siberian European EuropeaneSiberian EuropeaneSiberian European EuropeaneSiberian

MIS 3

LGT

BAIC

MUa

SU

NU

MU

SU

NU

MU

SU

NU

MU

SU

þ þ e þ e e þ þ þ þ þ þ þ þ e e e þ þ þ þ þ

þ þ e þ e e þ þ þ e þ þ þ e e e e þ þ þ þ þ

e þ e þ e e þ þ þ e þ þ e þ e þ e e þ þ e þ

e þ þ þ e þ þ þ þ e þ þ e þ e þ e e þ þ e þ

þ þ þ þ þ þ þ þ þ e þ þ þ þ þ e e e þ þ þ þ

e þ e þ e e þ þ þ e e e e þ e þ e e þ þ e þ

þ þ e þ e e þ þ þ e e e e þ e þ e e þ þ e þ

e þ e þ e þ þ þ þ e e e e þ þ e e e þ þ e þ

e þ e þ e e þ þ þ e e e e þ e þ e e þ þ e þ

þ þ e þ e e þ þ þ e e e e þ e þ e þ þ þ e þ

e þ e þ e þ þ þ þ e e e e þ þ e e e þ þ e þ

NU e Northern Urals, MU e Middle Urals, SU e Southern Urals.

Table 2 Species ratio of the EuropeaneSiberian, European and Siberian faunal assemblages in the carnivore fauna of the Urals during the Late Pleistocene (%%) Faunal complex

EuropeaneSiberian European Siberian Number of species a

NUa

MU

SU

MIS 5e

MIS 3

LGT

BAIC

MIS 5e

MIS 3

LGT

BAIC

MIS 5e

MIS 3

LGT

BAIC

e e e e

84 8 8 12

90 0 10 10

90 0 10 10

81 19 0 15

79 7 14 14

91 0 9 11

84 8 8 10

79 21 0 13

77 17 6 18

82 9 9 11

82 9 9 11

NU e Northern Urals, MU e Middle Urals, SU e Southern Urals.

Please cite this article in press as: Kosintsev, P.A., et al., Carnivores (Mammalia, Carnivora) of the Urals in the Late Pleistocene and Holocene, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.10.089

P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

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EuropeaneSiberian and Siberian faunal complex, while the fauna of the Middle and Southern Urals contained those from all the three faunal complexes (Table 2). At the end of this period, cave lion (Panthera spelaea) went extinct. The Early Holocene faunas have been studied in the Middle and Southern Urals (Table 3). The Early Holocene faunal composition of the Middle Urals and that of forest and steppe zones of the Southern Urals included the species of the EuropeaneSiberian, Siberian, and European faunal complexes (Table 4). The species from the EuropeaneSiberian complex dominated the faunas of all the areas. Range expansion resulted in reappearance of the otter (Lutra lutra) and European mink (Mustela lutreola) in the Uralian fauna. In the Early Holocene the Arctic fox (Vulpes lagopus) range began to decrease northwards, while the range of the steppe polecat (Mustela eversmanii) southwards. The Arctic fox disappeared from the fauna of the Southern Urals, but it persisted in the Middle Urals. The remnant population of the steppe polecat survived (Table 3).

reduced and they did not inhabit the Urals later (Table 1). Two other species became extinct. After the Last Glacial Maximum, the number of carnivores decreased. In the Late Glacial Transition (LGT) the fauna of the Northern Urals contained 10 species. There were 11 species in the Middle and the Southern Urals during this period (Table 1). The Late Glacial Transition fauna did not contain otter (Lutra lutra) and European mink (Mustela lutreola) whose ranges contracted. The faunas of the Northern and Middle Urals included the species of the EuropeaneSiberian and Siberian faunal complexes, while the Southern Urals contained the species of all the three complexes (Table 2). While sable, which is assigned to the Siberian faunal complex, inhabited the Northern and Middle Urals, the Southern Urals was distinguished by the presence of pine marten, which is assigned to the European faunal complex, and corsac, part of the Siberian faunal complex (Table 2).

Table 3 Species composition of carnivore fauna from the Urals during the Holocene and modern Species

NUa

Faunal complex

MU

SU “Mountain”

Lynx lynx Canis lupus Vulpes corsac Vulpes vulpes Ursus arctos Lutra lutra Gulo gulo Martes foina Martes martes Martes zibellina Meles anakuma Meles meles Mustela erminea Mustela eversmanii Mustela lutreola Mustela nivalis Mustela putorius Mustela sibirica a b

EuropeaneSiberian EuropeaneSiberian Siberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian EuropeaneSiberian European Siberian Siberian European EuropeaneSiberian EuropeaneSiberian European EuropeaneSiberian European Siberian

“Peneplain”

EHb

MH

LH

R

MH

LH

R

EH

MH

LH

R

EH

MH

LH

R

þ þ e þ þ þ þ e e þ e þ þ þ þ þ e e

þ þ e þ þ þ þ e e þ e þ þ e þ þ e e

þ þ e þ þ þ þ e þ þ þ e þ e þ þ e e

þ þ e þ þ þ þ e þ þ þ e þ e þ þ e þ

þ þ e þ þ þ þ e þ þ e þ þ e þ þ þ e

þ þ e þ þ þ þ e þ þ þ e þ e þ þ þ e

þ þ e þ þ þ þ e þ þ þ e þ e þ þ þ þ

þ þ þ þ þ þ þ e þ e e þ þ þ þ þ e e

þ þ þ þ þ þ þ e þ e e þ þ þ þ þ þ e

þ þ e þ þ þ þ þ þ e þ e þ þ þ þ þ e

þ þ e þ þ þ e e þ e þ e þ þ þ þ þ þ

e þ þ þ þ þ e e e e e þ þ þ þ þ e e

e þ þ þ þ þ þ e þ e e þ þ þ þ þ e e

e þ þ þ þ þ e e þ e þ e þ þ þ þ e e

e þ þ þ þ þ e e e e þ e þ þ þ þ e þ

NU e Northern Urals, MU e Middle Urals, SU e Southern Urals. EH e Early Holocene, MH e Middle Holocene, LH e Late Holocene, R e recent.

Table 4 Species ratio of the EuropeaneSiberian, European and Siberian faunal assemblages in the carnivore fauna of the Urals during the Holocene and modern (%%) Faunal complex

NUa

MU

SU «Mountain»

EuropeaneSiberian European Siberian Number of species a b

«Peneplain»

MHb

LH

R

EH

MH

LH

R

EH

MH

LH

R

EH

MH

LH

R

75 17 8 12

66 17 17 12

62 15 23 13

77 15 8 13

58 33 9 12

61 23 16 13

58 21 21 14

69 23 8 13

64 29 7 14

71 22 7 14

64 22 14 14

70 20 10 10

73 18 9 11

70 10 20 10

64 9 27 11

NU e Northern Urals, MU e Middle Urals, SU e Southern Urals. EH e Early Holocene, MH e Middle Holocene, LH e Late Holocene, R e recent.

During the Bølling e Allerød interstadial (BAIC), the faunas of the Northern and Southern Urals remained unchanged in comparison with those of the LGT (Table 1). Certain changes occurred in the faunal composition of the Middle Urals. It contained European badger (Meles meles). As a result, the faunal composition of the Northern Urals consisted of the species from the

During the Middle Holocene, the species of the EuropeaneSiberian faunal complex dominated the fauna of all the areas, but its share was noticeably smaller in the Middle Urals, than in the Northern and Southern Urals (Tables 3 and 4). The species of the European faunal complex closely follow those of the EuropeaneSiberian complex. As for the Siberian complex, it was represented

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P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

by one species only in all the areas, the sable (Martes zibellina) in the Northern and Middle Urals, and corsac (Vulpes corsac) in the Southern Urals. The Arctic fox (Vulpes lagopus) suffered further range contraction and eventually disappeared from the Middle Uralian fauna, though its remnant population still survived in the Northern Urals. Remnant population of the corsac (Vulpes corsac) still persisted in the forest zone of the Southern Urals. Range expansion led to appearance of the polecat (Mustela putorius) in the fauna of the Middle Urals and the forest zone of the Southern Urals. At the same time, the pine marten (Martes martes) range started to spread fast and led to its appearance in the Middle Urals (Gasilin et al., 2014). Only during the Middle Holocene, wolverine (Gulo gulo) appeared in the faunal composition of the steppe zone of the Southern Urals (Table 3). In the Late Holocene, the species of all three faunal complexes were represented in the faunas of all areas. The species of the EuropeaneSiberian complex dominated all the faunas (Table 3). The Siberian and European faunal complexes were represented by 1e4 species (Table 4). This period witnessed the end of a long process of range contraction of the steppe and tundra species. Thus, the Arctic fox (Vulpes lagopus) disappeared from the Northern Urals, while corsac (Vulpes corsac) dropped out of the forest zone of the Southern Urals (Table 3). Wolverine (Gulo gulo) withdrew from the steppe zone of the Southern Urals. Beach marten (Mates foina) appeared in fauna of forest part of Southern Urals only in the Late Holocene (Kosintsev, Gasilin, 2011; Gimranov, Kosintsev, 2015). At the onset of the Late Holocene, the Asian badger (Meles leucurus) forced out European badger (Meles meles) from the entire area of the Urals (Gasilin, Kosintsev, 2010). The transition to modern fauna of the Uralian carnivores (Table 3) has taken place over the last 200 years. At this time the Siberian weasel (Mustela sibirica) appeared in the Urals. Northward range extension of the pine marten (Martes martes) led to its appearance in the Northern Urals. At the same time wolverine (Gulo gulo) and Beach marten (Mates foina) disappeared from the forest zone of the Southern Urals (Table 3). Modern fauna is still characterized by the dominance of the species from the EuropeaneSiberian faunal complex, though the share of the species from the Siberian faunal complex has increased (Table 4). The analysis revealed that the carnivore faunal composition of the Urals in general and of its separate areas in particular underwent changes through the Late Pleistocene (Table 1) and Holocene (Table 3). The numbers of species and the shares of species assigned to EuropeaneSiberian, European, and Siberian faunal complexes changed in the faunal composition (Tables 2 and 4). The «megainterstadial» of the Last Glaciation (MIS 3) is characterized by the most prominent diversification of species, while in the Late Glacial Transition and the Bølling e Allerød interstadial the lowest number of species was recorded (Table 1). Species from the EuropeaneSiberian faunal complex dominated the faunas through the entire the Late Pleistocene and Holocene. The Late Pleistocene and Holocene show a general trend of fauna changes to a decreasing percentage of species from the European faunal complex and an increasing percentage of those from the Siberian faunal complex (Tables 2 and 4). Changes in the carnivore faunal composition resulted from several processes. Eight species (Crocuta crocuta spelaea, Cuon alpinus, Vulpes corsac, V. lagopus, Ursus thibetanus, Gulo gulo, Meles meles, Mustela eversmanii) suffered from range contraction. Today, the ranges of four species (Crocuta crocuta spelaea, Cuon alpinus Ursus thibetanus Meles meles) are beyond the Urals completely. Four species (Martes martes, Meles leucurus, Mustela putorius, Mustela sibirica) expanded their ranges. Four species (Lynx lynx, Lutra lutra, Martes foina, Mustela lutreola) inhabited the Urals in some periods, but were absent in others. Thus, their ranges fluctuated. Five

species (Canis lupus, Vulpes vulpes, Ursus arctos, Mustela erminea, Mustela nivalis) appeared consistently in the faunas of all the periods of study. One species (Panthera spelaea) was found in all the Late Pleistocene faunas, whereas two species (Ursus spelaeus, Ursus savini) persisted in the Uralian fauna only till the middle stage of the Late Pleistocene. The last three species went extinct. Thus, 16 out of 24 species underwent range shifts during the Late Pleistocene and Holocene. Range dynamics is apparently the principal cause of fauna changes, though species extinction also contributed to this process. These were exactly the same reasons for alterations of carnivore faunas in Europe (Sommer, Benecke, 2004, 2005; Turner, 2009; Stuart, 2014). Considering the carnivore distribution in the Urals in the Late Pleistocene and Holocene, the range limits of certain species here were a bit further north than in other areas. Here, we found for the Eem interglacial period the northernmost localities of Ursus thibetanus, U. savini, Meles meles (Makhnevskaya Ledyanaya cave, 59
260 N, 57
410 E), L. lynx (Bobylek grotto, 56
230 N, 57
370 E). There are for the second half of the MIS 2 (Late Glacial Transition and the Bølling e Allerød interstadial) the northernmost localities of Ursus spelaeus and U. savini (Medvezh'ya cave, 62
000 N, 58
44 E), Crocuta crocuta spelaea (Makhnevskaya cave, 59
260 N, 57
410 E), Martes zibellina (Tain cave, 59
250 N, 57
380 E), Mustela lutreola (Viasher cave, 59
050 N, 57
390 E), Meles meles (Viasher cave, 59
050 N, 57
390 E), Vulpes corsac (Georgievskaya cave, 57
020 N, 59
340 E), Cuon alpinus (Ignatievskaya cave, 54
540 N, 57
470 E) dating the MIS 3 (Fig. 1). In the Urals there are northernmost localities of Mustela eversmanii (Medvezh'ya cave, 62
000 N, 58
440 E) and M. meles (Lun'evka III cave, 59
100 N, 57
360 E) (Fig. 1.). The eastern limit of the U. spelaeus range lay in the Urals in the Late Pleistocene: its remains were found only on the western slope of the Urals and were absent on the eastern slope. In the Urals are the extreme north-eastern Holocene localities of M. foina (Podpornaya cave, 52
570 N, 55
590 E), M. eversmanii (Koksharovsko-Yurinskaya, 1, 58
100 N, 61
200 E) and M. lutreola (Kaninskaya cave, 62
020 N, 58
100 E) (Fig. 1). The time of extinction or range shifts of isolated species can be determined by radiocarbon dates obtained from their bones (Table A.2.). The available radiocarbon dates allow us to determine the time of the cave hyena (Crocuta crocuta spelaea) range contraction. Its bones gave 13 AMS dates (Table A.2.). 6 out of 13 dates are infinite. The latest dates were obtained from the bones of the Makhnevskaya cave (37700 ± 360, OxA-17003) in the Middle Urals and Ust-Katavskaya cave (35650 ± 450, OxA-10890) in the Southern Urals (Stuart, Lister, 2014). These dates are more ancient than those obtained from the localities situated in Central and Southern Europe and Britain (Stuart, Lister, 2014). This gives us good evidence to assume that the cave hyena's range contracted in the Urals earlier than in Europe. The carnivore fauna of the Urals is distinguished by three species that went extinct in the Late Pleistocene, namely Ursus spelaeus, Ursus savini and Panthera spelaea. The cave lion (Panthera spelaea) gave 11 AMS dates. The latest dates were obtained from the bones of the Pymva-Shor 2 locality (12995 ± 90, Ua-14006) in the Northern Urals and localities Podzemnyskh okhotnikov (13500 ± 65, OxA-11349), Verkhnegubakhinskaya cave (13560 ± 70, OxA-10909) and Viasher cave (13570 ± 70, OxA-10908) in the Middle Urals (Table A.2.). These dates are close to the dates obtained from the cave lion bones from the localities of France, Yakutia and Alaska (Stuart and Lister, 2011). These data show that the extinction of the cave lion in the Urals was synchronous with its extinction in northern Eurasia and North America. The cave bear bones provided 16 AMS dates (Table A.2.). The latest dates were obtained from the bones from the Tayn cave (37190 ± 680 BP) and Geologov 3 cave (38480 ± 360, OxA-19669) in the Middle Urals and

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P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

from the Asha 1 cave (47100 ± 900, OxA-16959) in the Southern Urals. These dates are more ancient than those obtained from the cave bear bones from Western Europe (Pacher and Stuart, 2008). The cave bear apparently went extinct in the Urals earlier than in Western Europe. Bones of the third extinct species, small cave bear Ursus savini, yielded only a few radiocarbon dates (Table A.2.). Of them, two AMS dates (31870 ± 190, OxA-16960; 36390 ± 270, OxA16964) were obtained from the bones from the Kizelovskaya cave in the Middle Urals. These dates allow us to infer that the small cave bear became extinct in the Urals at the end of Bryansk Interstadial (Middle Weichselian, MIS 3). Generally, extinction of the carnivores in the Urals was synchronous with their extinction in northern Eurasia (Stuart, 2014).

5. Conclusions We have studied the carnivore assemblages from the Late Pleistocene and Holocene localities of the Northern, Middle and Southern Urals. Analysis of the species composition of the carnivore fauna was performed for seven chronological periods: MIS 5e, MIS 3, mid-MIS 2 (LGT), late MIS 2 (BAIC), early MIS 1 (PreborealBoreal), mid-MIS 1 (Atlantic-Subboreal), late MIS 1 (Subatlantic). The MIS3 fauna consisted of 12 species in the Northern Urals, 14 species in the Middle Urals and 18 species in the Southern Urals. The Late Glacial Transition and the Bølling e Allerød interstadial faunas included 10e11 species. The faunal diversity and numbers increased in the Holocene: from 10 to 15 species inhabited different Uralian areas through different Holocene periods. Faunal composition comprised the species from European, Siberian and EuropeaneSiberian faunal complexes throughout all the Late Pleistocene and Holocene periods. Throughout all the periods it was dominated by the species from EuropeaneSiberian faunal complexes. The general trend of fauna changes is represented by a decreasing percentage of species from the European faunal complex and an increasing percentage of those from the Siberian faunal complex. Faunal composition has changed due to two main reasons, range shifts and extinction. Sixteen out of 24 species that inhabited the Urals during the Late Pleistocene and Holocene underwent range shifts. Their ranges shifted in different directions. Eight species (Crocuta crocuta spelaea, Cuon alpinus, Vulpes corsac, V. lagopus, Ursus thibetanus, Gulo gulo, Meles meles, Mustela eversmanii) suffered from range contraction. Four (Crocuta crocuta spelaea, Cuon alpinus, Ursus thibetanus, Meles meles) are no longer found in the Urals. The ranges contracted during the Late Pleistocene (Crocuta crocuta spelaea, Cuon alpinus, Ursus thibetanus) and Holocene (Vulpes corsac, V. lagopus, Gulo gulo, Meles meles, Mustela

7

eversmanii). Ranges of four species (Martes martes, Meles leucurus, Mustela putorius, Mustela sibirica) expanded. Three out of these four species (Meles leucurus, Mustela putorius, Mustela sibirica) appeared in the Urals for the first time. Range expansion occurred in the Holocene. Four species (Lynx lynx, Lutra lutra, Martes foina, Mustela lutreola) exhibited range fluctuations: while their ranges expanded during some periods and they were present in the Uralian fauna, during other periods their ranges contracted and they disappeared from the faunal composition. These fluctuations occurred both in the Pleistocene and Holocene. Five species (Canis lupus, Vulpes vulpes, Ursus arctos, Mustela erminea, Mustela nivalis) did not exhibit any range shifts throughout Late Pleistocene and Holocene. Three more species (Panthera spelaea, Ursus spelaeus, Ursus savini) did not undergo any range shifts, but these species went extinct during the Late Pleistocene. Two (Ursus spelaeus, Ursus savini) became extinct at the end of MIS 3 and one (Panthera spelaea) at the end of MIS 2. Because the presence of some species in the Urals was restricted to certain periods or certain areas, these species lend themselves to biostratigraphy and relative dating of the deposits. The remains of the Asiatic black bear (Ursus thibetanus), dhole (Cuon alpinus), cave bear (Ursus spelaeus), small cave bear (Ursus savini), cave hyena (Crocuta crocuta spelaea) and cave lion (Panthera spelaea) can be used for relative dating of the Late Pleistocene deposits all over the Urals. The remains of the polecat (Mustela putorius) are useful for relative dating of the Middle and Late Holocene, while those of the Asian badger (Meles leucurus) for relative dating of the Late Holocene all over the Urals. The remains of the corsac (Vulpes corsac), Arctic fox (Vulpes lagopus), wolverine (Gulo gulo), beech marten (Martes foina), pine marten (Martes martes), European badger (Meles meles) and steppe polecat (Mustela eversmanii) are appropriate for relative dating of the deposits from different Holocene periods in some Uralian areas. The history of the carnivore fauna in the Urals can be divided into the following principal stages: the MIS 5e fauna, the MIS 3 fauna, the fauna of the second half of the MIS 2, the MIS 1 fauna, and modern fauna. The changes in the carnivore fauna of the Urals, as well as of other European and Asian areas, occurred through the Late Pleistocene and Holocene to the present day. Acknowledgements The research was supported by the Russian Foundation for Basic Research (project N14-04-31231; 14-04-92104; 15-04-03882). Appendix

Table A.1 Carnivore species composition and remains numbers from the main localities in the Urals Species

Northern Urals MIS3

Lynx lynx Panthera spelaea Crocuta crocuta spelaea Canis lupus Cuon alpinus Vulpes corsac Vulpes lagopus Vulpes vulpes Ursus arctos

Mid-MIS 2

Late MIS2

Mid-MIS 1

1*

2

3

4

5

6

7

8

9

10

11

Late MIS 1 12

13

14

15

e 3 e e e e e e 1

e e e 79 e e 513 50 338

e e e 52 e e 125 e 62

e 2 e 4 e e 92 e e

1 e 27 e e 297 8 3

e 6 e 208 e e 1003 10 e

e 7 e 103 e e 103 1 3

2 e e 14 e e 20 5 1210

e e e 2 e e 2 37 40

e e e 1 e e 7 19 43

1 e e 1 e e 5 2 e

e e e 16 e e 3 e e

3 e e 13 e e 12 6 1128

e e e e e e e 10 61

1 e e 1 e e e 9 2

(continued on next page)

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P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

Table A.1 (continued ) Species

Northern Urals MIS3

Ursus tibetanus Ursus spelaeus Ursus savinii Lutra lutra Gulo gulo Martes foina Martes martes Martes zibellina Meles leucurus Meles meles M. erminea M. eversmanni M. lutreola M. nivalis M. putorius Species

Mid-MIS 2

Late MIS2

Mid-MIS 1

1*

2

3

4

5

6

7

8

9

10

11

12

13

14

15

e 253 1 e e e e e e e e e e e e

e e e e e e e 1 e e 28 18 e 26 e

e e e e e e e 4 e e 3 1 e 1 e

e e e e e e e

e e e

e e e e 11 e e 9 e e 32 21 e 18 e

e e e e e e e 6 e e e 2 e 2 e

e e e 16 33 e e 61 e 6 e e 2 e e

e e e e e e e 16 e e 1 e e e e

e e e 12 e e e 62 e 7 9 6 e 13 e

e e e 3 e e e 53 e 1 4

e e e 39 26 e e e e e e e e e e

e e e 43 76 e e 77 2 e e e e e e

e e e 13 e e e 99 e e e e 1 e e

e e e 6 2 e e 146 e e 34 e e 49 e

e e e 20

e e 16 e e 15 e

86 2 e 49 e

Late MIS 1

e 14 e

Middle Urals MIS 5e

MIS3

Mid-MIS 2

Late MIS2

Early MIS 1

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

Lynx lynx Panthera spelaea Crocuta crocuta spelaea Canis lupus Cuon alpinus Vulpes corsac Vulpes lagopus Vulpes vulpes Ursus arctos Ursus tibetanus Ursus spelaeus Ursus savinii Lutra lutra Gulo gulo Martes foina Martes martes Martes zibellina Meles leucurus Meles meles M. erminea M. eversmanni M. lutreola M. nivalis M. putorius

e 1 e 7 e e 1 3 2 1 23 41 e e e e e e 8 e e e e e

e e e 20 e e 47 14 e e 955 e e 1 e e 21 e 13 e e e e e

e 2 2 30 e e 27 9 e e 1080 e e e e e 3 e e 1 3 e 1 e

e e e 4 e e 5 e e e 24 e e e e e 2 e e e e e e e

1 1 e 13 e e 123 e 4 e e e e e e e 2 e e 88 e e 200 e

e 10 e 17 e e 44 5 92 e e e e e e e 7 e e e 2 e e e

e e e 8 e e 15 7 7 e e e e 2 e e e e e e e e e e

e e e 11 e e 24 11 15 e e e e 1 e e e e e e 3 e e e

2 e e 39 e e 104 7 2 e e e e 3 e e 11 e e 14 e e 1 e

e e e 34 e e 162 29 2 e e e e 9

e e e e e e 8 e 4 e e e e e e e e e 2 2 e e e e

e e e 2 e e 64 1 1 e e e e e e e 4 e e 5 5 e 5 e

e e e e e e 4 2 1 e e e e e e e 1 e e 9 e e 43 e

1 e e e e e 8 1 2 e e e e e e e 228 e e e e e e e

e e e 1 e e 2 7 e e e e e e e e 23 e e 14 e e 17 e

e e e 5 e e 12 4 11 e e e e 1 e e e e 1 5 e e 4 e

e e e 21 e e e e 5 e e e e e e e 4 e 8 e 1 e e e

e e e 20 e 1 8 e e e e e e 3 e e 6 e 1 10

e e e 1 e e 16 5 2

Species

Middle Urals

Lynx lynx Panthera spelaea Crocuta crocuta spelaea Canis lupus Cuon alpinus Vulpes corsac Vulpes lagopus Vulpes vulpes Ursus arctos Ursus tibetanus Ursus spelaeus Ursus savinii Lutra lutra Gulo gulo Martes foina Martes martes Martes zibellina Meles leucurus Meles meles M. erminea M. eversmanni

8 e e 4 4 e 1 e

e e 1 e e e 1 e 1 1 2 e 2 e

2 e

Sothern Urals

Mid-MIS 1

Late MIS 1

MIS 5e

MIS3

Mid-MIS 1

Late MIS 1

35

36

37

38

39

40

41

42

Mid-MIS 2 43

44

45

46

47

48

49

50

e e e 1 e e e e 13 e e e 5 e e 1 41 e 1 e e

e e e 2 e e e 3 8 e e e e 1 e 5 21 e e 2

1 e e 5 e e e 62 11 e e e 7 5 e e 105 e e 1 e

2 e e 26 e e e 15 95 e e e 11 1 e 13 1201 4 e 4 e

1 e e 3 e e e 7 e e 457 16 e e e e e e 6 1 e

e 5 28 55 e e 28 35 3 e 54 e 1 2 e 1 e e e e e

e 4 4 61 e e 30 11 e e 343 e e e e 2 e e e 2 e

e e 45 18 e e 5 6 9 e e e e e e e e e e e e

e 2 e 11 e 4 18 1 1 e e e e 1 e

e e e 2 1 e 7 6 e e e e e e e e e e e 1 2

e e e 1 e e

e e e

e e e 2 e 9 18 39 2 e e e e 1 e e e e e e 1

1 e e 1 e e e 3 e e e e e e e 1 e e e 67 1

e e e 11 e e e 19 26 e e e 6 e e 24 e 2 e 86 18

e e e e e e e 1 79 e e e 3 e 1 27 e e e e e

e e e 3 e

1 e e e e e e e 1 e e e 21 e

e 1 4 2 3 e e e e e e e e e e 24 1

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P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

9

Table A.1 (continued ) Species

M. lutreola M. nivalis M. putorius

Middle Urals

Sothern Urals

Mid-MIS 1

Late MIS 1

MIS 5e

MIS3

Mid-MIS 1

Late MIS 1

35

36

37

38

39

40

41

42

Mid-MIS 2 43

44

45

46

47

48

49

50

e e e

e e e

e e 1

e 1 e

1 1 e

e e e

e e e

e e e

e 1 e

e 6 e

e 23 e

e 4 e

e 1 e

e 38 e

1 9 e

e 1 e

*1 e Medvezhaya cave (inside gallery); 2 e Zhilische Sokola (pit 2); 3 e Usoltsevskaya cave; 4 e Cheremukhovo 1-1 (strata 9e12); 5 e Cheremukhova 1e4; 6 e Medvezhaya cave (strata grayish-brown “B” loamy soil); 7 e Medvezhaya cave (strata grey loamy soil); 8 e Kaninskaya cave (layer of Bronze age); 9 e Burmantovo (pit 1); 10 e Lisiy; 11 e Cheremukhovo 1-1 (strata 6); 12 e Uninskaya cave; 13 e Kaninskaya cave (layer of Iron age); 14 e Kamen’ Pysannyi; 15 e Cheremukhovo 1-1 (strata 1e4); 16 e Makhnevskaya-Ledyanaya cave; 17 e Bobylek (srata 5e6); 18 e Grotto Bolshoj Glukhoj (strata 9); 19 e Grotto Bolshoj Glukhoj (strata 8); 20 e Rasik; 21 e Verkhnegubakhinskaya cave; 22 e Bezymaynnyj; 23 e Zotinskij; 24 e Bobylek (strata 4); 25 e Pershinskaya cave; 26 e Lun'evka III cave; 27 e Dyrovatyj-na-Chusovoj; 28 e Lobvinskaya cave; 29 e Grotto Bolshoy Glukhoy (strata 2); 30 e Lobvinskaya cave; 31 e Dyrovatyj-na-Serge; 32 e Koksharovsko-Ur'inskaya; 33 e Bobylek (strata 3); 34 e Ust’-Koyvinskaya; 35 e Shigirskoe gorodische; 36 e Bobylek (strata 2); 37 e Turistov cave; 38 e Bobylek (strata 2); 39 e Barsuchyi dol; 40 e Ust’-Katavskaya cave; 41 e Ignat'evskaya cave; 42 e Smelovskaya cave; 43 e Nikol'skaya cave; 44 e Ignat'evskaya cave; 45 e Bayslan-Tash (strata 4); 46 e Balatukay; 47 e Syrtinskaya cave (strata 3); 48 e BayslanTash (strata 3); 49 e Bayslan-Tash (strata 2); 50 e Podpornaya cave.

Appendix

Table A.2 Radiocarbon dates for Quaternary carnivore remains from the localities of the Urals Species

Date(a)

±; (>) infinite dates

Lab. No (b)

Lat
N

Long
E

Localities

Dated object

Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Panthera spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Crocuta crocuta spelaea Canis lupus Canis lupus Vulpes corsac Vulpes lagopus Ursus savini Ursus savini Ursus savini Ursus savini Ursus savini Ursus savini Ursus savini Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus

12995 54500 29120 30140 >49600 13500 13570 13560 14750 41900 >39800 37700 >47600 35650 >59200 >40100 40240 44300 47500 >62300 51400 46200 >59800 >60800 40035 13675 25220 10180 >37890 31870 33980 36390 >48500 22750 26320 28390 >48600 >16470 26980 37190 39190 39580 39630 40340 47600 19550 22650 27070 27580 38480

90 2600 230 240

Ua-14006 OxA-10907 OxA-10894 OxA-10895 OxA-10888 OxA-11349 OxA-10908 OxA-10909 OxA-10910 OxA-10887 OxA-10845 OxA-17003 OxA-10889 OxA-10890 OxA-19533 OxA-10846 OxA-19660 OxA-19659 OxA-19537 OxA-19658 OxA-19536 OxA-17004 OxA-19535 OxA-19534 TUa-3525 AAR11385 GrA-41244 GrA-42216 SOAN-4528 OxA-16960 LE-2334 OxA-16964 no.? LU-3714 GIN-14244 SOAN-4799 no.? SOAN-4516 SOAN-4527 no.? OxA-16961 OxA-16965 OxA-16962 OxA-16963 OxA-16958 SOAN-4526 SOAN-4515 SOAN-4517 SOAN-4518 OxA-19669

67,10 60,42 60,24 60,24 59,25 59,17 59,05 58,53 58,40 54,54 54,10 59,26 55,11 54,56 54,56 54,54 54,54 54,54 54,54 54,54 54,51 53,20 53,20 53,20 66,34 59,00 57,02 64,55 59,25 59,06 59,06 59,06 59,06 54,33 54,33 62,05 62,05 59,25 59,25 59,25 59,25 59,25 59,25 59,25 59,25 59,06 59,06 58,46 58,46 58,46

60,51 60,22 60,03 60,03 57,46 57,50 57,40 57,38 57,34 57,47 56,51 57,41 58,38 58,08 58,08 57,47 57,47 57,47 57,47 57,47 59,58 59,00 59,00 59,00 62,25 58,00 59,34 51,44 57,46 57,65 57,65 57,65 57,65 57,16 57,16 58,05 58,05 57,46 57,46 57,46 57,46 57,46 57,46 57,46 57,46 57,65 57,65 57,43 57,43 57,43

Pymva-Shor 2 Shaytanskaya cave Cheremuchovo 1-1 cave Cheremuchovo 1e4 cave Tayn cave Podzemnykh okhotnikov Viasher cave Verkhnegubakhinskaya cave Kholodniy grotto Ignatievskaya cave Pobeda cave Makhnevskaya cave Sikiyaz-Tamak 7 Ust’-Katavskaya cave Ust’-Katavskaya cave Ignatievskaya cave Ignatievskaya cave Ignatievskaya cave Ignatievskaya cave Ignatievskaya cave Ustinovo grotto Smelovskaya 2 cave Smelovskaya 2 cave Smelovskaya 2 cave Mamontovaya Kur'ya Caves from r. Kos' va Georgievskaya cave Pizhma 1 Tayn cave Kizelovskaya cave Kizelovskaya cave Kizelovskaya cave Kizelovskaya cave Verkhnyaya cave Imanaj cave Medvezh'ya cave Medvezh'ya cave Tayn cave Tayn cave Tayn cave Tayn cave Tayn cave Tayn cave Tayn cave Tayn cave Viasher cave Viasher cave Geologov 3 cave Geologov 3 cave Geologov 3 cave

Bone Metapodia Metapodia Dentes Pelvis Calcaneus Vertebrae Scapula Bone Dentes Bone Mandibula Dentes Bone Dentes Bone Mandibula Dentes Dentes Cranium Dentes Dentes Dentes Dentes Bone Radius Pelvis Calcaneus Bone Mandibula Bone Cranium Cranium Bone Vertebrae, humerus Femur Bone Humerus Femur Bone Cranium Cranium Cranium Cranium Mandibula Bone Humerus Femur Femur Mandibula

65 70 70 70 1200 360 450

380 600 900 1500 1500

1825 70 130/120 50/45 1200 190 400 270 1210 1790 890 560 710 680 360 360 360 370 900 230 670 810 950 360

(continued on next page)

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10

P.A. Kosintsev et al. / Quaternary International xxx (2015) 1e11

Table A.2 (continued ) Species

Date(a)

Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus spelaeus Ursus arctos Ursus arctos Ursus arctos Ursus arctos Ursus arctos Ursus arctos Ursus arctos Martes zibellina Meles meles Meles meles Meles meles

>64400 29040 47100 >55500 >61500 >35900 44050 >27500 >27500 28700 37250 >46600 >50200 >62400 27500 47600 60600 12045 29527 2046 27100 8350 15840 2816 5600 7359 11340 11570

±; (>) infinite dates 385 900

1050

350 900 3600 79 320 33 250 750 70/60 35 70 79 50 50

Lab. No (b)

Lat
N

Long
E

Localities

Dated object

no.? SOAN-7043 OxA-16959 OxA-19666 OxA-19667 AA-90659 no.? IPAE-25 IEMEZ-723 LU-3715 LU-3876 LU-5135 LU-5134 OxA-19670 SOAN-5145 OxA-19679 OxA-19678 NSKA-AA-847 NSKA-AA-846 NSKA-848 SOAN-7916 OxA-no.? GrA-39266 no.? Ua-15104 AA-102248 GrA-39271 GrA-39270

58,16 55,00 55,00 55,00 55,00 54,55 54,55 54,54 54,54 54,33 54,33 54,33 54,33 54,33 54,10 54,10 54,10 62,05 62,05 62,00 61,55 60,43 57,27 54,54 67,10 62,00 59,10 59,05

57,59 57,18 57,18 57,18 57,18 57,53 57,53 57,47 57,47 57,16 57,16 57,16 57,16 57,16 56,51 56,51 56,51 58,05 58,05 58,09 60,05 60,17 61,27 57,47 60,51 58,09 57,55 57,39

Bolshoi Glukhoy Asha 1 Asha 1 Asha 1 Asha 1 Serpievskaya 2 cave Serpievskaya 2 cave Ignatievskaya cave Ignatievskaya cave Zapovednaya cave Zapovednaya cave Zapovednaya cave Zapovednaya cave Zapovednaya cave Pobeda cave Pobeda cave Pobeda cave Medvezh'ya cave Medvezh'ya cave Kaninskaya cave Severnaya cave Lakseyskaya cave Pershinskaya cave Zhemchuzhnaya cave Pymva-Shor 1 Kaninskaya cave Lunievka III Viasher cave

Bone Bone Mandibula Dentes Metacarpus Bone Bone Bone Bone Bone Bone Bone Bone Bone Bone Metatarsus Metacarpus Phalanx 2 Humerus Mandibula Bone Mandibula Ulna Bone Bone Mandibula Mandibula Bone

a

Dates are not calibrated. Laboratory codes for dates: AA e University of Arizona, Tucson, USA; AAR e University of Aarhus AMS Laboratory; GIN e Geological Institute, Russian Academy of Sciences (RAS); GrA e Groningen Rad. Lab., The Netherlands; IEMEG e Institute of Animal Evolution Morphology and Ecology, RAS; IPAE e Institute of Plant and animal Ecology, Ural division of RAS; LE e Institute of Archeology, Leningrad Branch (currently Institute of the History of Material Culture), RAS; LU e St. Petersburg Univ., RAS; NSKA e AMS pretreatment laboratory Centre of Cenozoic Geochronology Institute of Archaeology & Ethnography Siberian Branch of Russian Academy of Sciences; OxA e Oxford Accelerated, University of Oxford, UK; SOAN e Institute of Geology and Geophysics, Siberian Branch, RAS; TUa e Prepared in Trondheim, measured in Uppsala Univ., Sweden; Ua e Laboratory University of Uppsala, Sweden. b

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